Two new species of Actinocephalus (Eriocaulaceae) from Minas Gerais, Brazil

26 Accepted by M. Christenhusz: 8 Jul. 2011; published: 19 Aug. 2011

PHYTOTAXAISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2011 Magnolia Press

Phytotaxa 27: 26–36 (2011) www.mapress.com/phytotaxa/ Article

Two new species of Actinocephalus (Eriocaulaceae) from Minas Gerais, Brazil

LIVIA ECHTERNACHT1,2, MARCELO TROVÓ3 & PAULO TAKEO SANO1

1Laboratório de Sistemática Vegetal, Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, Rua do Matão 277, 05508-900, São Paulo, Brasil. [email protected], [email protected] 2UMR 7207 CNRS MNHN UPMC, Centre de Recherche en Paléobiodiversité et Paléoenvironnements, MNHN, 57 rue Cuvier, CC48, F-75005 – Paris, France3Evolution and Biodiversity of Plants, Faculty for Biology and Biotechnology. Ruhr University, D-44780 Bochum, Germany. [email protected]

Abstract

We describe and illustrate two new species, Actinocephalus delicatus and A. giuliettiae (Eriocaulaceae,Paepalanthoideae), from the Espinhaço Range in Minas Gerais, Brazil, and compare them with the morphologically mostsimilar species. Diagnostic characters, morphological variation, geographic distribution, habitat and conservation status,as well as line drawings, photos and a distribution map are provided for both species.

Resumo

Descrevemos e ilustramos duas novas espécies, Actinocephalus delicatus e A. giuliettiae (Eriocaulaceae,Paepalanthoideae), ocorrentes na Cadeia do Espinhaço, em Minas Gerais, Brasil, e as comparamos com as espéciesmorfologicamente mais similares. Discutimos caracteres diagnósticos, variações morfológicas, distribuição geográfica,habitat e status de conservação para ambas as espécies, além de apresentarmos ilustrações, fotos e um mapa dedistribuição geográfica.

Key words: Atlantic Forest, campo rupestre, Espinhaço Range, taxonomy

Introduction

In his Flora Brasiliensis, Koernicke (1863: 321) proposed Paepalanthus subg. Actinocephallus Koern., asubgenus later elevated to generic status by Sano (2004: 99). The genus was then characterized by paracladesof umbellate capitula and pistillate flowers with completely fused stigmatic branches. Costa (2005) proposedto enlarge this circumscription to include Paepalanthus subsect. Aphorocaulon Ruhland (1903: 167) inActinocephalus, to establish monophyly for the taxa as a result of several phylogenetic hypotheses (Giulietti etal. 2000, Unwin 2004, Andrade et al. 2010, Trovó 2010). Therefore, in this broad circumscription the genus ischaracterized by the presence of paraclades and simple (completely fused) stigmatic branche.

Actinocephalus is endemic to Brazil and can be found nearly throughout the country. Its area of highestdiversity lies in the “campos rupestres” (a Brazilian biome composed of rocky outcrops intermixed withgrasslands) in eastern Brazil, in a mountain range named Espinhaço, located in the states of Minas Gerais andBahia. Most species are endemic to the Espinhaço Range and several of them are microendemic, restricted toa few mountains (Giulietti & Hensold 1990, Costa et al. 2008).

Recent taxonomic revisions improved our understanding of particular taxa (Parra et al. 2010, Trovó &Sano 2010a, Costa 2005, Sano 2004, Parra 2000, Tissot-Squalli 1997, Hensold 1988), facilitating the

Phytotaxa 27 © 2011 Magnolia Press • 27NEW SPECIES OF ACTINOCEPHALUS (ERIOCAULACEAE)

recognition of undescribed species. Recent studies have earlier resulted in the description of new species of Eriocaulaceae from the Espinhaço Range (Echternacht & Sano in press, Trovó & Sano in press, Trovó et al.2011, Trovó & Sano 2010b). We therefore describe the following two microendemic species from the Espinhaço Range in Minas Gerais as new.

Taxonomy

Actinocephalus delicatus Sano, sp. nov. (Figs. 1, 2)Ab Actinocephalo brachypo habitu rhizomatoso, foliis glabris, scapis pilosis, bracteis paracladiorum intus glabris,

bracteis florum apicibus et sepalorum apicibus obtusis, bracteis involucralibus hialine usque ad brunnea et floribus pistillatis pedicellatis differt.

Type:—BRAZIL. Minas Gerais: Felício dos Santos, APA Felício dos Santos, Cachoeira do Sumidouro, 1350 m, 18o

13’S, 43o 15’W. 19 June 2006, P.L. Viana, F.S.F. Leite, L.E. Lopes & M. Ferreira 2512 (holotype SPF, isotype BHCB).

Perennial herbs, 10–18 cm tall, with a rhizome producing basal rosettes, and paraclades axillary to the rosette leaves with an umbel of inflorescences at its apex. Rhizome 1–6 cm long. Leaves rosulate, flat, patent, linear, 3.0–6.5 × 0.1–0.2 cm, glabrous, apex acute, margin glabrous. Paraclades 1–5 per rosette, unramified, 6.5–16.5 × 0.3 cm, hirsute; paracladial bracts distributed along the paraclade and subtending the umbel of capitules, erect, linear, 1.0–2.5 × 0.1–0.2 cm, adaxial surface glabrous, abaxial surface glabrescent, apex acute, margins glabrescent, base semiamplexicaul. Spathes 0.3–0.5 cm long, glabrous, apex acute. Scapes 18–35 per paraclade, 1.5–2.0 cm long, tomentose with simple trichomes; capitula 2–4 mm in diameter, hemispheric; involucral bracts hyaline to light-brown, obovate, ca. 2 mm long, glabrous, apex obtuse, ciliate toward the apex; receptacle hemispheric, pubescent. Flowers 3-merous, ca. 14 per capitula: 10 staminates, 4 pistilates; floral bracts castaneous to golden-brown, oblong, concave, ca. 2 mm long, glabrous, apex obtuse, ciliate toward the apex. Staminate flowers ca. 2 mm; pedicel ca. 0.5 mm long, with long trichomes; sepals colored like floral bracts, fused at the base, obovate, concave, ca. 1 mm long, glabrous, apex obtuse, ciliate toward the apex; corolla tubular, membranaceous, hyaline, ca. 1.5 mm long, 3-lobed, glabrous; stamens ca. 2 mm long; pistillodes 3, papillose. Pistillate flowers ca. 2 mm; pedicel ca. 0.5 mm long, glabrous; sepals colored like floral bracts, obovate, concave, ca. 2 mm long, glabrous, apex obtuse, ciliate toward the apex; petals hyaline, oblong, ca. 1 mm long, glabrous, apex acute, ciliate toward the apex; gynoecium ca. 2 mm long, stigmatic branches completely fused, twice as long as the nectariferous branches; staminodes 3, scale-like. Fruit a loculicidal capsule.

Distribution and ecology:—Actinocephalus delicatus is terrestrial or rupicolous and occurs in open areas, growing over rocky to sandy soils of the northeastern Espinhaço Range in Minas Gerais, Brazil. The species is known to occur in two distinct localities (Fig. 3): (1) in the Cachoeira do Sumidouro, in the Felício dos Santos municipality, a place predicted to be included in the Rio Preto State Park, where only a few individuals were found; and (2) in the Serra Negra State Park, in the Itamarandiba municipality, where a population of hundreds of individuals occurs throughout the Serra Negra. The individuals in the Cachoeira do Sumidouro were found in the riverbed, suggesting an aquatic habitat. These specimens were collected in June, in the dry season, so that it is unlikely that terrestrial individuals were flooded. However, we went to the area in the same season and, despite our intense search in the river edges and all around, we were unsuccessful to recover the species. As aquatic Actinocephalus are unknown, we suggest that the species is terrestrial, but some seeds germinated accidentally in the dry riverbed, where they were able to survive for a short while, without forming a perennial population. In addition, all the individuals in the Serra Negra are terrestrial or rupiculous, occurring in dry conditions, on sandy, free-draining soil.

IUCN Red List category:—Actinocephalus delicatus is considered endangered (E) according to criteria B2a (IUCN 2008).

ECHTERNACHT ET AL.28 • Phytotaxa 27 © 2011 Magnolia Press

FIGURE 1. Actinocephalus delicatus. A. Habit. B. Young capitulum. C. Involucral bract abaxially. D. Floral bract of the staminate flower abaxially. E. Staminate flower immature. F. Staminate flower immature with sepals removed and corolla opened. G. Pistillate flower. H. Gynoecium. (Drawn from the holotype)


FIGURE 2. Photographs of Actinocephalus delicatus. A. Habit. B. Inflorescence and capitula. C. Habitat in the SerraNegra State Park. D. Population and habit, individuals with immature inflorescences (Photos: Claudio Nicoletti—A, Band D—and Livia Echternacht—C).


FIGURE 3. Distribution map of Actinocephalus delicatus (black squares) and Actinocephalus giuliettiae (black stars).


Etymology:—The epithet refers to the delicate habit of the species, with its membranaceous leaves, and small and narrow paraclades.

Notes:—Actinocephalus delicatus matches morphological architectural pattern I (sensu Oriani et al.2008) due to its paraclades arising directly from a short aerial stem. Of all species exhibiting this pattern, A. delicatus shares morphological similarities with A. brachypus (Bong.) Sano (2004: 101) and A. herzogii (Moldenke) Sano (2004: 103). From the other Actinocephalus species it can be easily differentiated by its linear leaves, hirsute paraclades, hyaline to light-brown involucral bracts, and pedicelate pistillate flowers.Actinocephalus brachypus is the species morphologically most similar to A. delicatus. It shows approximately the same size of leaves, paraclades, and paracladial bracts, spathes and scapes, as well as a similar capitulum diameter. These common features give them a similar general habit, however they are easily distinguished, particularly by indumentum characters. Actinocephalus delicatus is a rhizomatous herb (vs. rhizome absent), with glabrous leaves (vs. hispid and ciliate), hirsute paraclades (vs. glabrous), and glabrous adaxial surface of the paracladial bracts (vs. hispid). It exhibits floral bracts and sepals of both staminate and pistillate flowers with obtuse apex (vs. acute). These species are also distinguished by the hyaline to light-brown involucral bracts (vs. dark-brown) and pedicelate pistillate flower (vs. sessile) in A. delicatus. Actinocephalus brachypus occurs in campos rupestres on the Diamantina Plateau (Minas Gerais), on roadsides between Diamantina and Gouveia (Sano 1999). Until now, both species were considered allopatric.

Actinocephalus herzogii is divided into two varieties, distinguished mainly by overall size (Sano 1999). Individuals of Actinocephalus herzogii var. humilis (Sano) Sano (2004: 103) are distinctly smaller, and morphologically similar to A. delicatus. Both are rhizomatous herbs with linear leaves, hairy paraclades and scapes. They have approximately the same size of paraclades and spathes and pedicelate pistillate flowers, a very unusual characteristic in the genus. Actinocephalus delicatus is distinguished by its longer leaves (3.0–6.5 cm vs. 2.0–2.5 cm) with glabrous margins (vs. ciliate), linear erect paracladial bracts (vs. lanceolate patent), spathes with acute apices (vs. truncate), and glabrous sepals of the pistillate flowers (vs. hairy in abaxial surface base). Actinocephalus herzogii var. humilis occurs in campos rupestres on Chapada Diamantina (BA), in the vicinity of Piatã (Sano 1999), thus, these species occur allopatrically.

Paratypes:—BRAZIL. Minas Gerais: Itamarandiba. Parque Estadual da Serra Negra, 18o 00’ 20.2’’S, 42o

43’ 28.9’’W, 13 September 2006, R.C. Mota, A.P. Fontana & K.A. Brahin 3089 (BHCB); Parque Estadual da Serra Negra, 18o 00’ 31.7’’S, 42o 43’ 46.7’’W, 1477 m, April 2010, L. Echternacht, T. V. Bastos, T. E. Almeida & A. M. de O. Santos 2140 (BHCB, P, SPF).

Actinocephalus giuliettiae Sano, sp. nov. (Figs. 4, 5)Ab Actinocephalo stereophyllo foliis rosulae majoribus (usque ad 10.0 cm × 5.5 cm) et cum apicibus acutis (non

mucronulatis), ramus elongatis pluribus (usque ad 15 × 5), spathis apicibus obliquus (non truncatis) et apicibus bractearum involucralium obtusis (non acutis nec apiculatis) differt.

Type:—BRAZIL. Minas Gerais: Itabirito, Serra de Capanema, Mina de Capanema, no alto da serra, 1726 m, 20o13’09”S, 43o34’41”W, 12 March 2009, L. Echternacht, F.F. do Carmo & I.C. de Campos 1944 (holotype SPF, isotypes BHCB, P).

Perennial herbs, 40–80 cm tall. Rhizome 1–3 cm long. Rosette bearing several axillary elongated branches, which bears the paraclades. Leaves rosulate, flat, patent, glaucescent, linear to lanceolate, 3–10 × 0.2–0.5 cm, glabrescent on both surfaces, slightly ciliated with short (ca. 0.5 mm) and long (ca. 3.5 mm) simple trichomes, apex acute, bearded in the young leaves, then glabrescent. Elongated branches indeterminate, 3–15 per rhizome, erect to patent, unramified, except for the paraclades, 35–60 × 0.3–0.6 cm, with bracts deciduous at the base and persistent at the middle and at the apex; elongate branch leaves densely spirally inserted, flat, patent, lanceolate, 2–5 × 0.2–0.5 cm, glabrescent on both surfaces, densely ciliated, especially at the base, with short and long simple trichomes, base semiamplexicaul, apex acute. Paraclades axillary, sub-opposite, unramified, 2.0–4.0 × 0.2 cm, pubescent; paracladial bracts distributed along the paraclade, subtending the


umbel of capitules and also inside the umbel, delimitating groups of scapes, flat, patent, lanceolate, 1.3–2.0 × 0.3–0.2 cm, adaxial surface glabrescent, abaxial surface pubescent to glabrescent, base semiamplexicaul, apex acute, ciliated with short and long simple trichomes. Spathes 0.8–1.3 cm long, pubescent to glabrescent, apex acute, ciliated and membranaceous. Scapes 40–50 per paraclade, 2.5–4.0 cm long, pubescent with patent trichomes, glabrescent when senescent; capitula 2.5–5.5 mm diam., hemispheric; involucral bracts brown, broadly ovate to elliptic, 1.0–2.0 mm long, pubescent abaxially, glabrous adaxially, apex obtuse, ciliated; receptacle hemispheric, pubescent. Flowers 3-merous, ca. 30 per capitula, half staminates, half pistilates; floral bracts castaneous, oblong, concave, ca. 2 mm long, pilose abaxially near the apex, apex obtuse and ciliate. Staminate flowers ca. 2 mm; pedicel ca. 0.3 mm long, with long trichomes; sepals castaneous to brown, fused at the base, broadly obovate, concave, ca. 1.5 mm long, pilose abaxially near the apex, apex round to mucronate, ciliate; corolla tubular, fleshy at the base, membranaceous and hyaline at the apex, ca. 1.5 mm long, 3-lobed, pilose adaxially at both apex sides, apex acute, glabrate; stamens ca. 2 mm long; pistillodes 3, papillose and pigmented. Pistillate flowers ca. 2.5 mm, sessile; sepals castaneous, obovate, concave, ca. 2.5 mm, pilose abaxially near the apex, apex obtuse to cuspidate, ciliate; petals lightly pigmented at the base, cream toward the apex, oblong, ca. 2.5 mm long, glabrous, apex obtuse, membranaceous and hyaline, ciliated at both sides but glabrous at the apex; gynoecium ca. 2.5 mm long, stigmatic branches completely fused, twice as long as the nectariferous branches; staminodes 3, scale-like. Fruit a loculicidal capsule.

Distribution and ecology:—Actinocephalus giuliettiae is terrestrial and occurs in open areas, growing over rocky quartzitic to ferruginous soils in the southern Espinhaço Range, in a mountain complex known as ‘Quadrilátero Ferrífero’ (Ferriferous Quadrangle), in the state of Minas Gerais, Brazil (Fig. 3). The species is known exclusively from the Serra do Capanema, in the Itabirito municipality, with one single population composed of less than 20 individuals.

IUCN Red List category:—The single population known for Actinocephalus giuliettiae occurs on a mining property predicted to be exploited in the near future. Therefore, according to the IUCN (2008) criteria B2a and C2ai, this species should be classified as critically endangered (CE).

Etymology:—This species is dedicated to Dr. Ana Maria Giulietti Harley. Her passionate work produced numerous branches and countless fruits, as she guided tens of Brazilian taxonomists. She is a pioneer in the studies of Eriocaulaceae systematics in Brazil and inspired all of us with her contributions to the knowledge of the family.

Notes:—Actinocephalus giuliettiae possesses a rhizome that gives rise to rosettes and lateral elongated branches ramifying into paraclades. In Actinocephalus, only two other species share a similar architectural pattern, A. falcifolius (Koern.) Sano (2004: 102) and A. stereophyllus (Ruhland) Sano (2004: 105), and which is described by Oriani et al. (2008) as pattern III. Actinocephalus giuliettiae differs from A. stereophyllus and A. falcifolius by its more numerous elongated branches per rosette, with bracts persistent at the middle and apex, longer scapes, with patent trichomes in youth, and petals of the pistillate flowers with glabrous apex. The presence of paracladial bracts inside the umbel of capitula has never been described for Actinocephalus, and might also be considered a diagnostic character for A. giuliettiae. There should be further investigation, however, to confirm whether A. stereophyllus, A. falcifolius or other species in the genus do not possess this character as well.

Actinocephalus stereophyllus is the species most similar to A. giuliettiae. Both have elongated branches similar in size, leaves of elongated branches are similar in shape and size, paraclades similar in size and in number of scapes, and spathes similar in size. In addition, the paraclades of A. stereophyllus usually are alternate, but they may be sub-opposite, as in A. giuliettiae. Actinocephalus giuliettiae differs from A. stereophyllus by its rosette leaves (up to 10.0 cm vs. 5.5 cm) with acute apices (vs. mucronulate), spathe apices oblique (vs. truncate) and involucral bracts with obtuse apices (vs. acute to apiculate). Actinocephalus stereophyllus occurs in campos rupestres on the Diamantina Plateau (Minas Gerais), in Diamantina municipality (Sano 2004), so that both species occur allopatrically.


FIGURE 4. Actinocephalus giuliettiae. A. Habit. B. Capitulum. C. Involucral bract abaxially. D. Floral bract of the staminate flower abaxially. E. Staminate flower. F. Staminate flower with all sepals one petal and one stamen removed. G. Pistillate flower. H. Petal of the pistillate flower showing the membranaceous, hyaline and glabrous apex I. Gynoecium. (Drawn from the holotype).


FIGURE 5. Photographs of Actinocephalus giuliettiae. A. Habit. B, C. Elongated branches, inflorescences and capitula.D. Rosette and elongated branches. E. Details of the inflorescence and capitula. F. Habitat in the Serra do Capanema.(Photos: Livia Echternacht).


Actinocephalus falcifolius is also very similar to A. giuliettiae. Both species have rosette leaves of equal size and pilosity and elongated branches of similar size. However, A. giuliettiae possesses erect rosette leaves (vs. falciform), elongated non ramified (vs. ramified) branches, sub-opposite (vs. alternate) paraclades, erect to ascendant (vs. sinuate to arched), longer branch leaves (2.0–5.0 cm vs. 1.5–2.5 cm) and erect (vs. falciform), erect paraclades (vs. arched), paracladial bracts ciliated all along the margin with long trichomes (vs. ciliated only at the base) and longer scapes (up to 4.0 cm vs. 2.5 cm). Actinocephalus falciformis also occurs in the Iron Quadrangle and several populations are known to occur in the municipalities of Ouro Preto, Lavras Novas and Itabirito (Sano 1999). It was not found in the Serra de Capanema and both species are thus currently occurring allopatrically.

Acknowledgments

The authors like to thank AcademicEnglishSolutions.com for the English corrections; to Flávio Fonseca do Carmo, Tiago Vilaça Bastos, Alessandra Monteiro dos Santos, Pedro Lage Viana, Heitor Bispo and Wanderley Lopes for fieldwork facilities; to Klei Sousa for the drawings of Actinocephalus delicatus. Livia Echternacht is financially supported by CAPES and FAPESP (proc 08/54886-1), Marcelo Trovó by Alexander von Humboldt Foundation, and Paulo Takeo Sano by CNPq (Proc. 307100/2009-0).

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Two new species of Actinocephalus (Eriocaulaceae) from Minas Gerais, Brazil

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